Nordic Journal of Botany 000: 001–015, 2016 doi: 10.1111/njb.00854, ISSN 1756-1051 © 2016 The Authors. Nordic Journal of Botany © 2016 Nordic Society Oikos Subject Editor: John Parnell. Editor-in-Chief: Torbjörn Tyler. Accepted 24 July 2015 New species of Ophiopogon, Peliosanthes and Tupistra (Asparagaceae s.l.) in the flora of Vietnam Leonid V. Averyanov, Noriyuki Tanaka, Khang Sinh Nguyen, Ba Vuong Truong, Duc Trong Nghiem and Tien Hiep Nguyen L. V. Averyanov ([email protected]), Komarov Botanical Inst., Russian Academy of Science, St Petersburg, Prof. Popov Str. 2, RU-197376, Russia. – N. Tanaka, 98-11 Otsuka, Hachioji, JP-192-0352 Tokyo, Japan. – K. S. Nguyen, Inst. of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Ha Noi, Vietnam. KSN also at: Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sciences, CN-510650 Guangzhou, P. R. China. – B. V. Truong, Inst. of Tropical Biology, Dept of Biological resources, 85 Tran Quoc Toan St., Distr. 3, Hochiminh City, Vietnam. – D. T. Nghiem, Dept of Botany, Hanoi Univ. of Pharmacy, 13-15, Le Thanh Tong str., Hoan Kiem district, Hanoi, Vietnam. – T. H. Nguyen, Center for Plant Conservation, no. 25/32, lane 191, Lac Long Quan, Nghia Do, Cau Giay District, Ha Noi, Vietnam. Five new species named Peliosanthes aperta, P. elegans, P. kenhillii, Tupistra densiflora and T. patula are described and illus- trated. These species are very local in distribution and endemic to northern or southern Vietnam. Two other species, Ophiopogon ogisui and Peliosanthes griffithii, are recorded as new to the flora of Vietnam. A key to the species of Tupistra occurring in Indochina and its neighboring regions is also provided. This paper focuses on the three generaOphiopogon Ker Gawl., conducted taxonomical and phytogeographical surveys of Peliosanthes Andrews (including Lourya Baill., Neolourya them in Vietnam and its adjacent regions in order to more L. Rodrig.) and Tupistra Ker Gawl. (for the generic closely elucidate their diversity (Tanaka 1998a, 1999a, circumscription see Tanaka 2010a) occurring in Vietnam. In 1999b, 1999c, 2000, 2001, 2003, 2004a, 2004b, 2010a, ‘Flore Générale de l’Indo-Chine’ (Vol. 6, fascicles 5 and 6 2010b, Averyanov 2011, Averyanov and Tanaka 2012, 2013, edited by Lecomte et al. 1934), Ophiopogon and Peliosan- Averyanov et al. 2013, 2014, 2015). thes (Neolourya) were placed in the family Haemodoraceae According to Tanaka, Tupistra consists of 20 species (Rodriguez 1934a), while Tupistra (as Gonioscypha) was in total (2010a), and Ophiopogon distributed in south classified under the Liliaceae (Gagnepain 1934a). In more and southeast Asia comprises 16 species (2001; 1 species, recent taxonomic treatises based on morphological and/or O. japonicus, is excluded, since it is not distributed spontane- molecular studies, they have been placed in the Asparagaceae ously here). Peliosanthes was reviewed by Jessop (1976) who s.l. (Tanaka 2010a, 2010b, Averyanov and Tanaka 2012, concluded that it comprises only 1 species, P. teta Andrews. 2013, Averyanov et al. 2013, 2014, 2015) or in the narrower In this treatment many other congeners previously published family Ruscaceae s.l. (Kim et al. 2010) or Convallariaceae as distinct were reduced to synonyms of P. teta. The only (Dahlgren et al. 1985, Conran and Tamura 1998, Tanaka character he found to differ among plants ofPeliosanthes was 1999a, Yamashita and Tamura 2000). The Asparagaceae the number of flowers in the axils of each bract. Based on s.l. as circumscribed by the Angiosperm Phylogeny Group differences in this character and in distribution he distin- (2003, 2009) is very broad, comprising several groups that guished two subspecies of P. teta. The taxonomic conclusion were formerly regarded as distinct families, like Ruscaceae reached by Jessop is thus remarkably different from that (Convallariaceae), Aphyllanthaceae and Agavaceae. These of previous botanists. In order to see how his study was families were treated as subfamilies under Asparagaceae by carried out, we reviewed his paper especially as to the meth- Chase et al. (2009), and the three genera in the present odology he employed. Jessop based his study on herbarium paper belong to the subfamily Nolinoideae that replaced the specimens, and he did not mention much about the mor- Ruscaceae. phological details of the flowers. Flowers of different taxa Members of the three genera distributed in Vietnam of Peliosanthes resemble each other in their basic structure, have hitherto been studied by botanists like Decaisne (1867), and they are mostly small and fleshy, so it is often difficult Gagnepain (1934a, 1934b), Rodriguez (1928, 1934a, to fully grasp floral features from dried and strongly pressed 1934b), Larsen (1961) and Jessop (1976). We have also specimens. Furthermore, herbarium specimens with flowers Early View (EV): 1-EV are generally not amply available and often lack informa- herein with taxonomic remarks and records. We also provide tion about characters like floral color, as Jessop himself a key to the species of Tupistra occurring in Indochina and stated. These factors are likely to have affected his conclu- its neighboring regions. sion, because they usually obscure specific delimitations that might exist. Jessop (1976) further demonstrated that several quantitative characters like the leaf length and leaf width of Material and methods the samples showed a continuous variation pattern. However, that result may be obtained in any plant group if quantita- In order to accurately identify our material, herbarium tive characters are sampled collectively from many specimens specimens (including type material when available) of many of a genus including multiple species. In our studies based known taxa of Ophiopogon, Peliosanthes and Tupistra kept at on living plants and/or herbarium specimens of Peliosanthes various herbaria were examined in advance. Surveys of these (Tanaka 2004a, 2004b, Averyanov and Tanaka 2012, 2013, specimens form a basis of this and preceding studies (Tanaka Averyanov et al. 2013, 2014, 2015), quite a few entities that 1998a, 1999a, 1999b, 1999c, 2000, 2001, 2004a, 2004b, appear distinct from other congeners were recognized. The 2010a, 2010b). degree of diversification into species in Peliosanthes appears In circumscribing species, we followed the following not much different from that in other plant groups. species concept (definition): “a species is a group of individu- Indochina is very rich in species diversity, at least of als similar in attributes and differing distinctly in taxonomi- several genera of Asparagaceae s.l. In our previous surveys, cally important characters from other such groups” (Tanaka we found as many as 22 new species of the three genera in 1996), or in short: “species are groups of similar individuals eastern Indochina (Vietnam, Laos and Cambodia), as listed differing distinctly from other such groups” (Tanaka 1998b). below (Tanaka 1999a, 2000, 2004b, 2010b, Averyanov These definitions are complemented by the following 2011, Averyanov and Tanaka 2012, 2013, Averyanov et al. explanation: ‘similar’ must be due to homology, and 2013, 2014, 2015, Vislobokov et al. 2014). ‘taxonomically important characters’ should be considered for each particular case. We believe that they accord well Ophiopogon fruticulosus Aver., N. Tanaka & K. S. Nguyen with the so-called taxonomic species concept that appears O. petraeus Aver. & N. Tanaka to have been most widely accepted. In practice, all kinds of O. rupestris Aver. & N. Tanaka character states (or attributes) that (a group of) individuals O. tristylatus Aver., N. Tanaka & Luu possess were taken into account when delimiting taxa. O. vietnamensis N. Tanaka Flowers of the three genera are more or less fleshy, Peliosanthes argenteostriata Aver. & N. Tanaka and their structures are generally largely distorted in the process of making herbarium specimens. So observations P. cambodiana Aver. & N. Tanaka of flowers were made mostly on living plants in their P. densiflora Aver. & N. Tanaka habitats and/or in cultivation. Further, when possible, P. divaricatanthera N. Tanaka inflorescences and flowers were preserved in spirit for P. grandiflora Aver. & N. Tanaka subsequent studies. P. hexagona Aver., N. Tanaka & K. S. Nguyen Specimens of the new species we are to describe here were P. lucida Aver., N. Tanaka & K. S. Nguyen collected in the field in 2014. Inflorescences and flowers P. micrantha Aver. & N. Tanaka were fixed and preserved in 60–70% ethanol. Measurements P. nivea Aver. & N. Tanaka of floral parts for description were made on both living and P. nutans Aver. & N. Tanaka liquid-preserved materials. Fresh flowers or floral parts were P. retroflexa Aver. & N. Tanaka found to shrink up to ca 20–30% in size in the drying pro- P. subcoronata N. Tanaka cess of making herbarium specimens. This was taken into P. triandra Aver. & N. Tanaka account when dried herbarium specimens were identified. In Tupistra breviscapa Aver. & N. Tanaka describing quantitative characters, infrequent extreme values T. khangii Aver., N. Tanaka & Vislobokov (i.e. rarely occurring minimal and maximal values) of a varia- T. laotica N. Tanaka tion range are parenthesized respectively before and after a T. theana Aver. & N. Tanaka normal variation range. The total number of new species of the three genera in the Ophiopogon ogisui M. N. Tamura & J. M. Xu (2007, above list is: Peliosanthes 13, Ophiopogon 5, and Tupistra 4. p. 39, Fig. 1) (Fig. 1) Our field surveys in Indochina are still underway, and in our prospect, even more new species will be discovered. In this Type: China, Guangxi, Longzhu, Jinlong, Gaoshan, Banbi, respect, the current state of our knowledge of these genera in ca 440 m a.s.l., Mikinori Ogisu 250 (holotype: PE, n.v.). eastern Indochina is quite incomplete. In our latest survey in Vietnam, we discovered five addi- Habitat and phenology tional new species, three of which belong to Peliosanthes Shady primary and secondary lowland forests of evergreen (P. aperta, P. elegans and P. kenhillii) and two to Tupistra broad-leaved trees on limestone at elevations about 400– (T.